Greeting and Salutations Grand Practitioners of CS generation,

 Moving on now to fresh water animals. Once the ancestors of the modern freshwater animals had made the transition to the freshwater environment, presumably by way of the estuaries, there was no longer any great advantage to their descendants in continuing to maintain body fluids as concentrated as seawater, as long as they remained in their new environment. Such excessively hypertonic internal conditions simply aggravated the problems of obtaining enough salt and bailing out excess water. Thus it is understandable that natural selection should have favored a reduction of the osmotic concentration of the body fluids within the bounds possible for the continuance of the life of the tissues, and that modern freshwater animals, both invertebrates and vertebrate, have osmotic concentrations decidedly lower than seawater. It seems incompatible with cellular existence, however, for the body fluids to be as dilute as freshwater, for no organisms are actually isotonic with their freshwater medium; the body fluids of freshwater animals are typically hypotonic relative to seawater, but hypertonic relative to freshwater.
 Now, if freshwater animals are hypertonic relative to the surrounding environmental medium, there will be a strong tendency for water to move into the organism and for salts to be lost from the organism to the surrounding water. At first glance, the obvious evolutionary solution to this problem might seem to be the development of completely impermeable membranes covering the entire body, but further thought shows that this solution would have been impracticable, since a truly aquatic organism must maintain some permeable membranes exposed to the water for gas exchange. Because mammals that live in the water breath air and hence need never expose permeable respiratory membranes to the water, they can maintain an impermeable barrier between their body fluids and the water in which they live. But fully aquatic freshwater animals cannot use the  "method of evasion" exclusively.  They must also be able to carry out active osmoregulation, which usually involves excretory organs that can  pump out the water as fast as it floods in--- preferably through the production of urine more dilute than the body fluids--- and/or special secretory cells somewhere on the body that can absorb salts from the environment and release them into the blood. Both corrective measures--production of dilute urine and absorption of salts (minerals/metals) --entail movement of material against concentration gradients and therefore necessitate expenditure of energy.
 An examination of the water and salt regulation typical of modern freshwater bony fishes will provide a good example of the before mentioned processes. The bony fishes are the class Osteichthyes, most of the fish familiar to you. They have backbones of vertebra. The blood and tissue fluids of the fish are more concentrated than the environmental water. The method of evasion is used to the extent that much of the body is covered by relatively impermeable skin and scales, and that the fish almost never drink. There is, however, a constant osmotic intake of water across the membranes of the gills and of the mouth, and a constant loss of salts across the same membranes. The method of correction is used two ways: The excess water is eliminated in the form of very dilute and copious urine produced by the kidneys, and salts are actively absorbed by specialized cells in the gills.
 Curiously enough, marine vertebrates, bony fishes living in the seawater have the reverse problem: They live in water, yet they steadily lose water to their environment and are in constant danger of dehydration. The explanation is that the ancestors of the bony fishes (vertebra fishes) apparently lived in freshwater, not in the sea, and that when some of  their descendants moved to the marine environment they retained their dilute body fluids. Vertebra were a development of estuary animals, and moved back to the sea and on to the land. Thus marine bony fishes are hypotonic relative to the surrounding water, and they have the problem of excessive water loss and excessive salt intake. Besides benefiting from the evasive adaptation of relatively impermeable skin and scales, they use two corrective measures: They drink almost continuously to replace the water they are constantly losing, and, by means of specialized cells in the gills, they actively excrete the salts they unavoidably take in with the water. Most of the nitrogenous wastes are excreted as ammonia through the gills; hence only a small quantity of urine is produced by the kidneys, and little water need be lost in this manner. Apparently fish kidneys have not evolved the capacity to produce concentrated urine, and they are  consequently of no help in salt elimination.
 The marine elasmobranch fishes (sharks and their relatives) probably also evolved from freshwater ancestors, but they solved the osmotic problem in a very different way.  Their blood contains about the same concentrations of salts as the blood of marine bony fishes, but their blood also contains high concentrations of urea, to which they are more tolerant than most vertebrates. By conserving the urea instead of excreting it, the marine elasmobranchs maintain a total osmotic concentration in their blood slightly greater than that of the seawater.  They therefore have no problem of water loss. Excess salt is excreted by special glandular cells in the rectum.

    ---to be continued---
 

Bless you,   Bob Lee

--
oozing on the muggy shore of the gulf coast
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